Chelodina Longicollis (Shaw 1794) – Eastern Long-Necked Turtle, Common Long-Necked Turtle, Common Snake-Necked Turtle

Conservation Biology of Freshwater Turtles and Tortoises: A Compilation Project ofChelidae the IUCN/SSC — Chelodina Tortoise and longicollis Freshwater Turtle Specialist Group 031.1 A.G.J. Rhodin, P.C.H. Pritchard, P.P. van Dijk, R.A. Saumure, K.A. Buhlmann, J.B. Iverson, and R.A. Mittermeier, Eds. Chelonian Research Monographs (ISSN 1088-7105) No. 5, doi:10.3854/crm.5.031.longicollis.v1.2009 © 2009 by Chelonian Research Foundation • Published 13 November 2009

Chelodina longicollis (Shaw 1794) – Eastern Long-Necked Turtle, Common Long-Necked Turtle, Common Snake-Necked Turtle

Ro d Ke n n e t t 1, Jo h n Ro e 2, Ka t e Ho d g e s 2, a n d Ar t h u r Ge o r g e s 2

1North Australian Indigenous Land and Sea Management Alliance (NAILSMA), Charles Darwin University, Darwin NT 0909, Australia [[email protected]]; 2Institute for Applied Ecology, University of Canberra, ACT 2601, Australia [[email protected], [email protected], [email protected]]

Su m m a r y . – The eastern long-necked turtle, Chelodina longicollis (Family Chelidae), has a wide distribution throughout southeastern Australia. It occupies a broad range of freshwater aquatic habitats but is more abundant in shallow, ephemeral wetlands often remote from permanent rivers. Its propensity for long distance overland migration, coupled with a low rate of desiccation and the capacity to estivate on land, enable it to exploit highly-productive ephemeral habitats in the absence of competition from fish and other turtle species. In wetter periods, such habitats provide optimal conditions for growth and reproduction. In drier periods, however, turtles may need to seek refuge in permanent water where high population densities and low productivity can lead to reduced growth rates and reproductive output. The species is an opportunistic carnivore that feeds on a broad range of plankton, nekton and benthic macro-invertebrates, carrion, as well as terrestrial organisms that fall upon the water. It is relatively slow to mature (7–8 yrs for males and 10–12 yrs for females), lays between 6 and 23 hard-shelled eggs during spring and late summer, and can produce up to 3 clutches per year. Although currently considered common and not under major threat, the most widespread conservation concern for C. longicollis is high nest predation from the introduced fox (Vulpes vulpes), and roads, pest fencing, and habitat changes brought about by prolonged drought and climate change, which present localized and potential future threats for certain populations. Di s t r i b u t i o n . – Australia. Found throughout southeastern Australia, including southeastern Queensland, New South Wales, Victoria, and southeastern South Australia. Sy n o n y m y . – Testudo longicollis Shaw 1794, Emys longicollis, Chelodina longicollis, Hydraspis longicollis, Chelys (Chelodina) longicollis, Chelydura longicollis, Chelodina longicollis longicollis, Chelodina novaehollandiae Duméril and Bibron 1835, Chelodina sulcata Gray 1856a, Chelodina longicollis sulcata, Chelodina sulcifera Gray 1856b, Chelodina longicollis sulcifera. Su b s p e c i e s . – None currently recognized. St a t u s . – IUCN 2008 Red List: Not Listed (= Least Concern, LR/lc) (assessed 1996, needs updating); CITES: Not Listed; Australian EPBC Act: Not Listed.

Figure 1. Adult female Chelodina longicollis from Booderee National Park, Jervis Bay, New South Wales, Australia. Photo by John Roe. 031.2 Conservation Biology of Freshwater Turtles and Tortoises • Chelonian Research Monographs, No. 5

Figure 2. Dorsal and ventral view of the shell of a female Chelodina longicollis. Note the broadly expanded plastron, especially anteriorly, the darkening of the scute boundaries of the plastron, and the recessed intergular scute. Photos by Arthur Georges.

Taxonomy. — Chelodina longicollis was the first variation (Hodges, unpubl. data), and are not widely Australian freshwater turtle to be described, having been accepted. The taxonomy of the species is no longer regarded collected by Sir Joseph Banks during Captain Cook’s first as controversial. voyage down the east coast of Australia in 1770 (Cann 1998). The genus Chelodina comprises three distinct First described as Testudo longicollis by Shaw (1794), it was clades (Goode 1967; Burbidge et al. 1974, Georges and subsequently considered distinct enough to be placed in its Adams 1992: Seddon et al. 1997: Georges et al. 2002), own genus Chelodina by Fitzinger (1826). commonly regarded to be subgenera (Legler 1985). Chelodina sulcata, described by Gray (1856a) and Chelodina longicollis belongs to one of these clades then erroneously referred to as Chelodina sulcifera by (subgenus Chelodina), containing also C. steindachneri, Gray (1856b), was for a while recognized as distinct from C. canni, C. pritchardi, C. mccordi, C. reimanni, and C. C. longicollis (Müller 1936), but later shown to be a form novaeguineae. Chelodina longicollis is the sister taxon to of C. longicollis in which abnormal shell growth, possibly the remaining species in the clade (Georges et al. 2002). as a consequence of a dietary deficiency, resulted in the The species hybridizes freely in nature with both C. retention of impressions of earlier plastral scute growth canni and C. rugosa to produce viable offspring, but the (Budde 1968). processes that maintain genetic identity of the species are Several authors have noted morphological differences not known. among some populations of C. longicollis. Chessman Description. — The carapace is moderately broad, (1978) suggested that turtles east and west of the Great expands slightly posteriorly and has a pronounced medial Dividing Range might be accorded subspecific status, and groove on the 2nd to 4th vertebrals that deepens with Beck (1991) proposed eastern and southern races, but these age. The marginals immediately above the tail are raised suggestions have not been supported by rigorous analysis. medially and the lateral marginals tend to curl upwards in Neither arrangement is supported by mtDNA sequence older animals. The color of the carapace varies from black

Figure 3. Head of Chelodina longicollis from Eastlakes Swamps, Figure 4. Dorsal and ventral view of a hatchling Chelodina lon- Sydney, New South Wales. Photo by John Cann. gicollis. The distinctive bright orange coloration of the plastron fades with age. Photos by Oliver Römpp. Chelidae — Chelodina longicollis 031.3

Figure 5. Distribution of Chelodina longicollis in Australia. Red points = museum and literature occurrence records based on Iverson (1992) plus more recent and authors’ data; green shading = projected distribution based on GIS-defined hydrologic unit compartments (HUCs) constructed around verified localities and then adding HUCs that connect known point localities in the same watershed or phys- iographic region, and similar habitats and elevations as verified HUCs (Buhlmann et al., in press), and adjusted based on authors’ data. Populations on Bass Strait islands north of Tasmania are considered introduced and are not shown here. to light-brown, with the sutures outlined in black in lighter in dune lakes at Jervis Bay, NSW, they are 237 mm and colored specimens. The plastron is large, usually extending 200 mm (Kennett 1987); and in the Latrobe Valley they are laterally beyond the inner edges of the ventral surfaces of 216 mm and 188 mm (Chessman 1978). These differences the marginal scutes that border the axillary pocket, and likely reflect local habitat productivity as much as any has a deep anal notch. It is cream in color with the sutures broad latitudinal trends (Kennett 1987). broadly edged in black. The neck is long, approximately Hatchlings are almost entirely black, with a vivid 60% of the carapace length (CL) and covered with small orange stripe running along either side of the lower jaw pointed tubercles. Unlike the Chelodina within species and neck. Orange spots are present in the center of each group “B” (Goode 1967), the head and neck are relatively plastral shield and the underside of the marginal scutes narrow and, when withdrawn, are protected by both of the carapace. This vivid coloration may serve to warn carapace and plastron. The dorsal surfaces of the head, aquatic predators of the unpalatability of the hatchling neck, and limbs are dark gray to brown and the undersides (Dorrian and Ehmann unpubl. data). Hatchlings have a are cream to yellow. Both fore and hind limbs have four mass of 4–5 g and are ca. 30 mm in CL. claws and the tail is quite short. When disturbed, it releases Distribution. — Chelodina longicollis is distributed a pungent yellow fluid from inguinal and axillary scent throughout southeastern Australia, from the vicinity of glands that earns it the affectionate colloquial name of Port Lincoln on the Eyre Peninsula west of Adelaide in “stinker” (Eisner et al. 1978). South Australia, throughout the coastal rivers of Victoria Females tend to have deeper shells, grow to larger sizes and New South Wales (NSW) north to the Fitzroy River than males, and the ventral surface of the posterior lobe of drainage of coastal Queensland. It is found throughout the their plastron is generally roundly convex as opposed to Murray-Darling drainage, in the Paroo drainage and in the the concave or irregular shape of the males. The tail of the headwaters of the Cooper Creek drainage in inland eastern female tends to be shorter and fatter than the male but is Australia. Specimens have been recorded west of the Great usually withdrawn tightly into the shell when handled, thus Dividing Range as far north as the Burdekin River. the distinction is not always clear. Maximum adult body Early reports of C. longicollis from tropical northern sizes vary geographically. Maximum female and male CL Australia are probably Chelodina canni (Kennett et al. in riverine habitat in the Murray Valley are 282 mm and 1992). Where the species comes into contact with C. 249 mm, respectively (Chessman 1978); in the ponds of canni, in the coastal plains of the Styx River drainage in the Gungahlin suburb in Canberra, ACT, they are 279 mm Queensland, the two species freely hybridize (Georges et and 214 mm (Rees 2008); in farm dams near Armidale, al. 2002). Chelodina longicollis occurs in sympatry with NSW, they are 239 mm and 199 mm (Parmenter 1976); Emydura macquarii and Chelodina expansa throughout 031.4 Conservation Biology of Freshwater Turtles and Tortoises • Chelonian Research Monographs, No. 5

much of the southern portion of its range (Chessman 1978; Chessman 1988a) and with Elseya albagula, Myuchelys latisternum, Emydura krefftii, and C. expansa in the north (Legler and Cann 1980; Georges 1982b). There are apparently introduced populations on some of the islands in the Bass Strait north of Tasmania, specifically Chappell Island in the Flinders group (W. Austr. Mus. 12049) and King Island (Templeton 1972). Habitat and Ecology. — Chelodina longicollis is active and can forage at low temperatures (Chessman 1988b); consequently it is able to penetrate into higher latitudes and altitudes than other Australian turtles. The species utilizes a broad range of habitats throughout its range, including permanent riverine waterholes, lakes, farm dams, and shallow temporary ponds. However, it occurs in greatest abundance in shallow, ephemeral waterholes or Figure 6. Chelodina longicollis has a strong propensity for in bodies of water that are remote from permanent rivers, overland migration, and can spend extensive periods estivating in terrestrial refugia. This animal is buried in leaf litter in Booderee often in the absence of other turtle species (Chessman National Park. Photo by John Roe. 1988a). Primarily a bottom dwelling species, it is rarely seen swimming at the surface, but occasionally leaves the 1987; Roe et al. 2008). Overland migration enables C. water to bask (Webb 1978). It is most active in the early longicollis to exploit abundant food resources in temporary morning and late afternoon (Chessman 1988b). water bodies in the absence of competition from fish and Chelodina longicollis exhibits a suite of characteristics other species of turtle (Chessman 1984b; Georges et al. that enable it to exploit ephemeral and semi-permanent 1986; Chessman 1988a). At Jervis Bay, NSW, terrestrial waterholes, including (1) a marked propensity for overland migration is part of a dynamic pattern of habitat utilization travel and ability to navigate (Stott 1987; Graham et al. (Kennett 1987; Kennett and Georges 1990; Roe et al. 1996; Roe and Georges 2007); (2) a slow rate of desiccation 2009). In drier periods, most turtles are confined to the (Chessman 1984a; Roe et al., 2008); and an ability to refuge of permanent freshwater dune lakes where high (3) drink pooled water while terrestrial; (4) absorb free population densities coupled with low productivity result water through its cloaca; and (5) physiologically adjust in drastically reduced growth and reproduction. In wetter the composition of its urine to retain salts and reduce periods, turtles move out into ephemeral wetlands where water loss (Rogers 1966; Roe 2008). These behavioral conditions for growth and reproduction are extremely and physiological attributes allow C. longicollis to either favorable (Kennett and Georges 1990; Roe and Georges estivate on land to await the re-flooding of wetlands, or 2008a). Competitive exclusion by other turtle species and to travel overland to permanent water bodies (Chessman 1983; Roe and Georges 2008a, b). An estivation duration of up to 480 days, without returning to water, has been observed (Roe and Georges 2007); however, energy stores ultimately limit survival out of water where they cannot feed (Roe et al. 2008). Estivation sites are typically located under canopy in forested habitats where turtles bury in leaf litter several cm deep near woody structures such as shrubs and logs, maintaining lower shell temperatures than when active (Roe et al. 2008; Rees 2008). Terrestrial estivation can continue into winter, requiring turtles to hibernate on land. The migratory habits of C. longicollis are well known (Anonymous 1941; Stott 1987; Chessman 1988a; Beck 1991). The species is regularly encountered crossing roads that traverse wetland areas and colonizes farm dams within a few years of their construction (Parmenter 1976). In some cases overland migrations are extremely frequent, with some individuals (males, females, and Figure 7. The range of habitats occupied by Chelodina longicollis juveniles) associating with several different water bodies includes brackish waters. Increased salinization of Australia’s during a single year (Roe 2007; Rees 2008; Roe et al. freshwater rivers is a threat to freshwater turtles, and leads to novel 2009). Overland travel usually coincides with rainfall encounters, in this case, of a turtle and an encrusting estuarine tube worm in Lake Alexandrina, South Australia. This animal and treks covering several kilometers and more than 40 was still alive, but suffering restrictions on its mobility. Photo by days on land have been reported (Chessman 1978; Stott Sally Grundy. Chelidae — Chelodina longicollis 031.5 fish may explain its relatively scarcity in large permanent at Armidale (NSW) mature at 7 and 10–11 yrs respectively, waterholes elsewhere in its range (Chessman 1978; similar to the 8–12 yrs for turtles from the Murray River Georges et al. 1986). (Chessman 1978). Growth rates of juvenile turtles also Chelodina longicollis is an opportunistic carnivore vary geographically. Kennett and Georges (1990) recorded that feeds on a broad range of planktonic, nektonic, and average juvenile (CL = 80 mm) growth rates at Jervis Bay benthic macro-invertebrates, carrion, as well as terrestrial of 7 mm/yr, compared to around 22 mm/yr for similar sized organisms that fall upon the water (Parmenter 1976; juveniles in the Murray Valley (Chessman 1978). Mean Chessman 1984b; Georges et al. 1986). Turtles feed with adult growth rates are uniformly low, often less than 1 mm/ a “strike and gape” action in which prey are effectively yr, and many adults do not grow at all for long periods. For sucked into the mouth by a vacuum created when the example, mature females occupying a large semi-natural hyoid is lowered (Parmenter 1976). The composition waterhole near Canberra, showed no measurable increase of the diet varies little with the size or sex of the turtle, in size when re-measured 20 years after first capture except that adults tend to eat more carrion. Competition (Georges, unpubl. data). with fish may significantly reduce dietary intake because Estimates of longevity in natural populations are turtles occupying waterholes with fish had significantly rare given that turtles are difficult to age past the point at lower stomach content volumes than turtles in waterholes which they mature, and few long term capture-recapture without fish (Chessman 1984b). Turtles are not selective in studies have been undertaken for long enough to document the animals that they eat and geographic variation in their longevity. Parmenter (1985) speculated that, at an average diet probably results from differences in local abundances adult growth rate of 0.51 mm/yr, a recently matured of prey (Georges et al. 1986). female of 170 mm and 10 yrs of age would take 143 yrs Data on growth rates are limited to populations in to reach the maximum size of 240 mm. This is likely to the southerly portion of its range where turtles become be an overestimate because few turtles actually reach the inactive during cold winter months and feeding and growth maximum recorded size for a population, but it suggests cease (Chessman 1988b). Growth rates are also markedly that wild turtles are likely to live at least as long as captive influenced by food availability. Kennett and Georges animals (36 yrs, Goode 1967; 37+ yrs, Goin and Goin 1971 (1990) demonstrated significant differences between local in Parmenter 1976). In the longest reported continuous populations in both adult and juvenile growth rates that study of this species, several turtles captured as adults in correlated with measured differences in food availability 1983 were recaptured as late as 2007 in our Jervis Bay, between the habitats. Turtles in a highly productive NSW studies, indicating minimum ages of 31–34 yrs in the ephemeral swamp grew substantially faster than those in wild (Roe 2007). permanent dune lakes where food availability was lower. Adult survivorship in populations occupying perm- Growth rates of turtles in the lakes were faster in wetter anent water in farm dams is high (98% per year, Parmenter years when the area of useable habitat and presumably food 1985). Survivorship rates of turtles in the dune lakes of availability were greatest; whereas, many individuals, both Jervis Bay, NSW, ranged from 85% to 94% for both adults and juveniles, failed to grow at all during drier years. juveniles and adults (Roe et al. 2009), but lower survival Growth and adult body size differences were also observed rates (55%) were observed for turtles estivating near dry between turtles in suburban ponds and an adjacent nature ephemeral wetlands during a prolonged drought (Roe and reserve in the ACT. The faster growth rates and larger body Georges 2008a). Predators on adults include the White- sizes of suburban turtles were hypothesized to reflect high breasted Sea Eagle (Haliaeetus leucogaster), dingos nutrient runoff from golf courses, roads, and gardens, as (Canis familiaris) and foxes (Vulpes vulpes), and many well as the more stable water levels from urban runoff are killed by motor vehicles on roads or when entrapped that afforded suburban turtles a longer growing season by pest-proof fencing (J. Roe, pers. obs.). over those of the nature reserve, where wetlands dried for Mating usually occurs in September (Chessman 1978; several months of the year and forced the turtles to estivate Parmenter 1985; Kennett and Georges 1990) and lacks (Rees 2008; Rees et al. in press). the often elaborate courtship behaviors that occur in other Size at maturity varies geographically but shows no species (Murphy and Lamoureaux 1978). Nesting occurs clear latitudinal trends. The largest sizes at maturity are in the late spring and early summer (October – January) recorded in the Murray Valley (NSW and Victoria), where and up to three clutches containing between 6 and 23 eggs males mature at CL of 180–190 mm and females at 210 mm (average of 14) are laid (Vestjens 1969; Parmenter 1976, (Chessman 1978). Smaller sizes are reported from Jervis 1985; Chessman 1978; Legler 1985; Georges, unpubl. Bay (NSW), Armidale (NSW) and the Latrobe Valley data). Eggs are white, hard-shelled and ellipsoid in shape, (Victoria) where males mature at CL of 142–152 mm and about 20 mm wide and 30 mm long and weigh 6–7 g. females at 165–185 mm (Parmenter 1976; Chessman 1978; Females may travel up to 500 m or more from water to Kennett and Georges 1990). Age estimation is hampered preferred nesting sites on a crest or ridge at the top of a by erratic growth and the loss and non-annual formation of slope and will nest in a variety of substrates from soft sand scute annuli (Stott 1988), but the available data suggest that to hard clay or even gravel roads. Water is periodically age at maturity is less variable than size. Males and females released from the cloaca into the nest to soften the ground 031.6 Conservation Biology of Freshwater Turtles and Tortoises • Chelonian Research Monographs, No. 5

(Vestjens 1969) and one female was observed to return to been previously inaccessible due to physical barriers such the water several times to take on more water before she as extensive stretches of dry land (Beck 1991; Doody et. al. was able to complete nesting (Cann 1978). Once the eggs 2006; Rees et al. in press; Roe and Georges in press). have been laid the female pushes earth back into the hole Estimates of population densities range from 26 to with her rear feet and tamps it down by raising herself on 400 turtles/ha (Parmenter 1976; Chessman 1978; Georges her legs above the nest and dropping abruptly to compact 1982a). Large fluctuations in population size may result the earth. from climate-mediated mass immigration or emigrations Despite the female’s attempts to camouflage her nest, of turtles (Kennett and Georges 1990; Roe et al. 2009). predation on nests is high. Parmenter (1985) reported There are anecdotal reports of appreciable declines a minimum estimate of 49% of nests destroyed, Beck in populations in the Brisbane catchment of southeast (1991) and Thompson (1983) reported predation rates in Queensland over the past 20 years. This may be due to the excess of 90%. Predators include native animals such as combined factors of increased wetland loss and degradation water rats (Hydromys chrysogaster), goannas (Varanus associated with urban development, increased road sp.) and Australian ravens (Corvus coronoides) but mortality of migrating turtles and increased fox predation most nests (> 90%) are destroyed by the introduced fox (John Cann, pers. comm.). (Thompson 1983; Parmenter 1985; Beck 1991). Eggs Threats to Survival. — The introduced fox is the in nests that escape predation take between 110 and primary predator on nests and, in some years and locations, 150 days to complete incubation and hatchlings emerge every nest is destroyed. Turtle populations in habitats where in autumn, usually after rain has softened the ground fox predation is high are often dominated by larger (= older) (Parmenter 1985; Vestjens 1969). Overwintering in the adults and juveniles are absent (Thompson 1983). This lack nest by hatchlings has not been directly observed but may of successful nesting and juvenile recruitment has prompted occur in populations in Gippsland, Victoria (Parmenter concerns that the long term survival of many populations of 1985; Chessman 1978). C. longicollis is in jeopardy, but that in the short term, such Clutch size or number of clutches per year varies among population declines are masked by the longevity of adults populations but no clear latitudinal trends are apparent. (Thompson 1983; Thompson 1993a; Thompson 1993b). Larger females tend to lay larger clutches, hence estimates Foxes also kill adults and large winter kills of turtles by of clutch size will be influenced by the size distribution foxes have been observed (Georges, unpubl. data). of females sampled from a population. Reproductive Riverine habitat modification for agricultural ind- output also varies with annual and spatial variation in food ustries and urban development threaten populations of availability (Kennett and Georges 1990), further obscuring C. longicollis. Water has been extracted from Australia’s any geographical trends. Under good conditions all females Murray-Darling Basin, the primary inland habitat for C. in a population will nest but reproduction is depressed and longicollis, since the early 1900s, and approximately 75% may cease entirely in overcrowded conditions or where of the flow of water in the Murray-Darling Basin is now food is severely limited (Kennett and Georges 1990). redirected for human use. Changes in river flow, discharge Adult sex ratios in natural populations range from and flood frequency, the addition of physical barriers equal to both male and female biased (Parmenter 1976; to movement, increased river and dry land salinity, and Kennett and Georges 1990). Females in a crowded drought a decline in water and habitat quality due to prolonged refuge maintained better body condition (i.e., were heavier) drought and climate change all have negative impacts on than equivalent sized males, suggesting that female-biased the abundance and distribution of C. longicollis (Roe and sex ratios may result from male-biased mortality during Georges in press). Habitat modification especially affects times of severe food limitation (Kennett and Georges populations of C. longicollis, where productive semi- 1990). Rees (2008) did not observe skewed sex ratios in permanent wetlands are converted to permanent water areas bisected by suburban roads. Skewed adult sex ratios lakes such as occurred in Lake Mokoan in Victoria (Cann are unlikely to be the result of biased hatchling sex ratio 1993). Toxic algal blooms coupled with a reduced food because hatchling sex is genetically determined, and not availability in the lake led to the starvation and eventual temperature-dependent as in many other turtle species death of large numbers of turtles. (Georges 1988; Palmer-Allen et al. 1991). Migrating turtles travel long distances overland Population Status. — Chelodina longicollis is common (distances of 8 km have been reported) and such movements in all major river systems throughout its broad range. between habitats are an essential part of the species’ strategy While agricultural and urban development has had adverse of habitat use. Land use changes such as roads and fencing effects on the natural habitat of the species, this may have that impede the free travel of turtles may have a deleterious been offset by the construction of numerous artificial water impact on populations over a broad area. However, Rees bodies for agricultural and pastoral purposes (Parmenter (2008) concluded that populations in suburban Canberra 1976; Rennie 2002; Doody et. al. 2006). It is also likely were not severely threatened by roads and other urban that the provision of new habitats such as dams, river hazards. Most turtles could avoid crossing roads during impoundments, irrigation channels, and urban ponds have their normal movements by travelling through under-road facilitated population expansion into areas that may have stormwater drainage culverts (Rees et al. in press). Chelidae — Chelodina longicollis 031.7

Conservation Measures Taken. — The species is New South Wales. Surrey Beatty and Sons, Sydney, pp. 363–65. currently protected under both state and federal legislation Ca n n , J. 1998. Australian Freshwater Turtles. Sydney and which prohibits the taking and killing of native wildlife by Singapore: Beaumont Publishing, 292 pp. all except Aboriginal people and Torres Strait Islanders. Ch e s s m a n , B.C. 1978. Ecological studies of freshwater turtles in south-eastern Australia. Ph.D. Dissertation, Monash University, It is regarded as a common species by all States and Melbourne. Territories where it occurs, and individuals may be kept Ch e s s m a n , B.C. 1983. A note on aestivation in the Snake-necked without a permit provided they are not collected from Turtle, Chelodina longicollis (Shaw) (Testudines: Chelidae). the wild and are obtained from a legal source. Overseas Herpetofauna 14:96–97. export of native animals is regulated by the Environment Ch e s s m a n , B.C. 1984a. Evaporative water loss from three south- Protection and Biodiversity Act 1999 administered by the eastern Australian species of freshwater turtle. Australian Journal Australian government. of Zoology 32:649–655. Chelodina longicollis is largely ubiquitous within Ch e s s m a n , B.C. 1984b. Food of the Snake-Necked Turtle, Chelodina permanent and ephemeral freshwater and brackish wetlands longicollis (Shaw) (Testudines: Chelidae) in the Murray Valley, Victoria and New South Wales. Australian Wildlife Research of all varieties and so is found in all national parks and 11:573–578. protected areas within its occupied range, including the Ch e s s m a n , B.C. 1988a. Habitat preferences of freshwater turtles following National Parks (among many others): Booderee, in the Murray Valley, Victoria and New South Wales. Australian Jervis Bay, Seven Mile Beach, and Thirlmere Lakes in Wildlife Research 15:485–91. New South Wales. Ch e s s m a n , B.C. 1988b. Seasonal and diel activity of freshwater Conservation Measures Proposed. — Specific turtles in the Murray Valley, Victoria and New South Wales. conservation plans for C. longicollis are not a high priority Australian Wildlife Research 15:267–76. and have not been formulated, as the species is considered Do o d y , S., Os b o r n e , W., Bo u r n e , D., Re n n i e , B. a n d Si m s , R. common throughout its range. 2006. Vertebrate Biodiversity on Australian Rice Farms: An Inventory of Species, Variation Among Farms, and Proximate Captive Husbandry. — Chelodina longicollis are Factors Explaining that Variation. Rural Industries Research and hardy in captivity and thrive on a balanced diet of fish, Development Corporation, Canberra Australia. Publication No. tadpoles, invertebrates and prepared food. Eggs can be W05/198. Project No. UCA-6A. readily obtained by hormonal injections to induce egg- Du m é ri l , A.M.C. a n d Bibr o n , G. 1835. Erpetologie Generale ou laying. Histoire Naturelle Complete des Reptiles. Roret, Paris, 680 pp. Current Research. — Current mtDNA based studies Ei s n e r , T., Jo n e s , T.H., Me i n wa l d , J., a n d Le g l e r , J.M. 1978. of the phylogeography of C. longicollis are underway Chemical composition of the odorous secretion of the Australian (Hodges, unpubl. data) and the taxonomy and biogeography turtle, Chelodina longicollis. Copeia 1978:714–715. of the entire genus Chelodina is currently under review Fit z i n g e r , L.J. 1826. Neue classification der reptilien nach ihren naturlichen verwandtschaften. Vienna: Heubner. 66 pp. (Georges and Thomson, 2009). Ge o rg e s , A. 1982a. Diet of the Australian freshwater turtle Emydura krefftii (Chelonia: Chelidae), in an unproductive lentic Literature Cited environment. Copeia 1982:331–336. Ge o rg e s , A. 1982b. Ecological studies of Krefft’s River Tortoise, An o n y m o u s . 1941. Death on the plains-where instinct fails. Wild Emydura kreftii, from Fraser Island, Queensland. Ph.D. life (Melbourne.) 3:13–14. Dissertation, University of Queensland, Brisbane. Be c k , R.G. 1991. The Common Longnecked Tortoise Chelodina Ge o rg e s , A. 1988. Sex determination is independent of incubation longicollis (Shaw 1802) (Testudinata: Chelidae): A comparative temperature in another chelid turtle, (Chelodina longicollis). study of the morphology and field behaviour of disjunct Copeia 1988:248–254. populations. South Australian Naturalist 66:4–22. Ge o rg e s , A. a n d Ad a m s , M. 1992. A phylogeny for Australian Be y n o n , F. 1991. Comparative rates of oxygen consumption and chelid turtles based on allozyme electrophoresis. Australian development in two species of turtle from the genus Chelodina. Journal of Zoology 40:453–476. Honours Dissertation, University of Canberra, Canberra. Ge o rg e s A., Ad a m s M. a n d McCo r d , W. 2002. Electrophoretic Bu d d e , H. 1968. A rare Australian chelid? International Turtle and delineation of species boundaries within the genus Chelodina Tortoise Society Journal 2(5):12-15. (Testudines: Chelidae) of Australia, New Guinea and Indonesia. Bu h l m a n n , K.A., Ak r e , T.S., Iv e r s o n , J.B., Kara p ata k i s , D., Zoological Journal of the Linnean Society 134:401–421. Mitt e r m e i e r , R.A., Ge o rg e s , A., Rh o d i n , A.G.J., v a n D ij k , P.P., a n d Ge o rg e s A. a n d Th o m s o n S. 2009. Diversity of Australasian Gibb o n s , J.W. In press. A global analysis of tortoise and freshwater freshwater turtles, with an annotated synonymy and keys to turtle distributions with identification of priority conservation areas. species. Zootaxa, in review. Chelonian Conservation and Biology 8(2):in press. Ge o rg e s , A., No rri s , R. a n d We n s i n g , L. 1986. Diet of the Bu rbi d g e , A.A., Kir s c h , J.A.W. a n d Mai n , A.R. 1974. Relationships freshwater turtle Chelodina longicollis (Testudines: Chelidae), within the Chelidae (Testudines: Pleurodira) of Australia and from the coastal dune lakes of the Jervis Bay Territory. Australian New Zealand. Copeia 1974:392–409. Wildlife Research 13:301–308. Ca n n , J. 1978. Tortoises of Australia. First Edition. Angus and Go o d e , J. 1967. Freshwater tortoises of Australia and New Guinea. Robertson, Sydney. Lansdowne Press Pty Ltd, Melbourne. Ca n n , J. 1993. Don’t take our freshwater turtles for granted. In: Gra h a m T.E., Ge o rg e s A. a n d McEl h i n n e y , N. 1996. Terrestrial Lunney, D. and Ayers, D. (Eds.). Herpetology in Australia - A orientation by the eastern long-necked turtle, Chelodina longicollis, Diverse Discipline. Transactions of the Royal Zoological Society of from Australia. Journal of Herpetology 30:467–477. 031.8 Conservation Biology of Freshwater Turtles and Tortoises • Chelonian Research Monographs, No. 5

Gra y , J.E. 1856a. On some new species of freshwater tortoises for coping with wetland drying in freshwater turtles. Ecology from North America, Ceylon and Australia, in the collection of 89:485–494. the British Museum. Proceedings of the Zoological Society of Ro e , J.H. a n d Ge o rg e s , A. 2008b. Terrestrial activity, movements, London 1855[1856](23):197-202. and spatial ecology of an Australian freshwater turtle, Chelodina Gra y , J.E. 1856b [“1855”]. Catalogue of Shield Reptiles in the longicollis in a temporally dynamic wetland system. Austral Collection of the British Museum. Part I. Testudinata (Tortoises). Ecology 33:1045-1056. British Museum, London, 79 pp. Ro e , J.H. a n d Ge o rg e s , A. In press. Responses of freshwater turtles to Iv e r s o n , J.B. 1992. A Revised Checklist with Distribution Maps of the drought: the past, present and future implications of climate change Turtles of the World. Richmond, IN: Privately printed, 363 pp. in Australia. In Gow, K. and Paton, D. (Eds.). The Phoenix of Natural Ke n n e tt , R. 1987. Reproduction, growth and dispersal of the eastern Disasters: Impact of Drought. Nova Science Publishers, New York. long-necked turtle Chelodina longicollis Shaw (Testudinata: Ro e , J.H., Ge o rg e s , A. a n d Gr e e n , B. 2008. Energy and water flux Chelidae) in the Jervis Bay Territory’. Honours Dissertation, during terrestrial estivation and overland movement in a freshwater Australian National University, Canberra. turtle. Physiological and Biochemical Zoology 81:570-583. Ke n n e tt , R.M. a n d Ge o rg e s , A. 1990. Habitat utilization and its Ro e , J.H., Bri n t o n , A.C., Ge o rg e s , A. 2009. Temporal and spatial relationship to growth and reproduction of the eastern long- variation in landscape connectivity for a freshwater turtle in a necked turtle, Chelodina longicollis, (Testudinata: Chelidae), temporally dynamic wetland system. Ecological Applications from Australia. Herpetologica 46:22–33. 19:1288–1299. Ke n n e tt , R.M., Ge o rg e s , A., Th o m a s , K. a n d Ge o rg e s , T.C. 1992. Ro g e r s , L.J. 1966. The nitrogen excretion of Chelodina longicollis Distribution of the long-necked freshwater turtle Chelodina under conditions of hydration and dehydration. Comparative novaeguineae and new information on its ecology. Memoirs of Biochemistry and Physiology 18:249–260. the Queensland Museum 32:179–182. Se d d o n J.M., Ge o rg e s A., Ba v e r s t o c k P.R., a n d McCo r d W. Le g l e r , J. 1985. Australian chelid turtles: reproductive patterns 1997. Phylogenetic relationships of chelid turtles (Pleurodira: in wide-ranging taxa. In: Grigg, G., Shine, R. and Ehmann, Chelidae) based on mitochondrial 12S rRNA gene sequence H. (Eds.). Biology of Australasian Frogs and Reptiles. Royal variation. Molecular Phylogenetics and Evolution 7:55–61. Zoological Society of New South Wales, Sydney, pp. 117–123. Sh aw , G. 1794. The Zoology of New Holland. British Museum of Le g l e r , J.M. a n d Ca n n , J. 1980. A new genus and species of chelid Natural History, London. turtle from Queensland, Australia. Contributions to Science (Los St o tt , P. 1987. Terrestrial movements of the freshwater tortoise, Angeles County Museum of Natural History) 324:1–18. Chelodina longicollis (Shaw), as monitored with a spool tracking Mü l l e r , L. 1936. Über die Artberechtigung von Chelodina sulcifera device. Australian Wildlife Research 14:559–567. Gray. Zoologische Anzeiger, Leipzig 116:273-287. St o tt , P. 1988. Use of growth rings to determine age on the freshwater Mu r p h y , J.B. a n d La m o u r e a u x , W.E. 1978. Mating behaviour in tortoise Chelodina longicollis: a cautionary note. Transactions of three Australian chelid turtles (Testudines: Pleurodira: Chelidae). the Royal Society of South Australia 112: 179–180. Herpetologica 34:398–405. Te m p l e t o n , M.T. 1972. Reptiles of King Island. Tasmanian Naturalist Pa l m e r -Al l e n , M., Be y n o n , F. a n d Ge o rg e s , A. 1991. Hatchling 31:1-2. sex ratios are independent of temperature in field nests of Th o m p s o n , M.B. 1983. Populations of the Murray River Tortoise the long-necked turtle, Chelodina longicollis (Testudinata: Emydura (Chelidae): the effect of egg predation by the red fox, Chelidae). Wildlife Research 18:225–231. Vulpes vulpes. Australian Wildlife Research 10:363–371. Par m e n t e r , C.J. 1976. The natural history of the Australian freshwater Th o m p s o n , M.B. 1993a. How secure are “common tortoises”? turtle Chelodina longicollis Shaw (Testudinata: Chelidae). Ph.D. Australian Natural History 24:72. Dissertation, University of New England, Armidale. Th o m p s o n , M.B. 1993b. Hypothetical considerations of the Par m e n t e r , C.J. 1985. Reproduction and survivorship of Chelodina biomass of chelid tortoises in the River Murray and the possible longicollis (Testudinata: Chelidae). In: Grigg, G., Shine, R. and influences of predation by introduced foxes. In: Lunney, D. and Ehmann, H. (Eds.). Biology of Australasian Frogs and Reptiles. Ayers, D. (Eds.). Herpetology in Australia - A Diverse Discipline. Royal Society of New South Wales, Sydney, pp. 53–61. Transactions of the Royal Zoological Society of New South Re e s , M. 2008. Impacts of urbanisation on the eastern long-necked Wales. Surrey Beatty and Sons, Sydney, pp. 219–224. turtle (Chelodina longicollis). Honours Dissertation, University Ve s tj e n s , W.J.M. 1969. Nesting, egg-laying and hatching of the of Canberra, Canberra, ACT. snake-necked tortoise at Canberra, A.C.T. The Australian Re e s , M., Ro e , J.H., a n d Ge o rg e s , A. In press. Life in the suburbs: Zoologist 15:141–151. Behavior and survival of a freshwater turtle in response to drought We bb , G.J.W. 1978. Observations on basking in some Australian and urbanization. Biological Conservation. turtles (Reptilia: Testudines: Chelidae). Herpetologica 34:39–42. Re n n i e , B. A. 2002. Habitat use and movement patterns of the eastern long-necked turtle (Chelodina longicollis) in rice agroecosystems Citation Format for this Account: of N.S.W. Unpublished Honours Dissertation, University of Canberra, Canberra. Ke n n e tt , R., Ro e , J., Ho d g e s , K., a n d Ge o rg e s , A. 2009. Che- Ro e , J.H. 2007. The terrestrial ecology of a freshwater turtle, lodina longicollis (Shaw 1784) – eastern long-necked turtle, Chelodina longicollis, in Booderee National Park, Australia. Ph.D. common long-necked turtle, common snake-necked turtle. In: Dissertation, University of Canberra, Canberra, ACT. Rhodin, A.G.J., Pritchard, P.C.H., van Dijk, P.P., Saumure, Ro e , J.H. 2008. Chelodina longicollis (Eastern long-necked turtle). R.A., Buhlmann, K.A., Iverson, J.B., and Mittermeier, R.A. Drinking Behavior. Herpetological Review 39:212-213. (Eds.). Conservation Biology of Freshwater Turtles and Tor- Ro e , J.H. a n d G e o rg e s , A. 2007. Heterogeneous wetland complexes, toises: A Compilation Project of the IUCN/SSC Tortoise and buffer zones, and travel corridors: landscape management for Freshwater Turtle Specialist Group. Chelonian Research freshwater reptiles. Biological Conservation 135:67–76. Monographs No. 5, pp. 031.1-031.8, doi:10.3854/crm.5.031. Ro e , J.H. a n d G e o rg e s , A. 2008a. Maintenance of variable responses longicollis.v1.2009, http://www.iucn-tftsg.org/cbftt/.